Abstract
We describe two new species of skinks from Gunung Penrissen, Sarawak, Malaysia, in northern Borneo, Tytthoscincus batupanggah sp. nov. and T. leproauricularis sp. nov. Morphological and molecular analyses both corroborate the two new species as unique compared to all other Tytthoscincus and additional Sphenomorphus that are candidates for taxonomic placement in the genus Tytthoscincus. Despite their phenotypic similarity and sympatric distribution, a molecular analysis shows that the new species are not sister taxa and exhibit a deep genetic divergence between each of their respective sister taxa. We discuss how historical climatic and geographic processes may have led to the co-distribution of two relatively distantly related phenotypically similar species. In light of these discoveries, we also emphasize the importance of conserving primary montane tropical rainforest for maintaining species diversity.
References
Aljanabi, S.M. & Martinez, I. (1997) Universal and rapid salt-extraction of high quality genomic DNA for PCR-based techniques. Nucleic Acids Research, 25, 4692–4693.
http://dx.doi.org/10.1093/nar/25.22.4692Annandale, N. (1912) Zoological results of the Abor expedition, 1911–1912. Records of the Indian Museum, 8, 7–59.
http://dx.doi.org/10.5962/bhl.part.1186Arévalo, E., Davis, S.K. & Sites, J.W. (1994) Mitochondrial DNA sequence divergence and phylogenetic relationships among eight chromosome races of the Sceloporus grammicus complex (Phrynosomatidae) in central Mexico. Systematic Biology, 43, 387–418.
http://dx.doi.org/10.1093/sysbio/43.3.387Boulenger, G.A. (1894) A list of the reptiles and batrachians collected by Dr. E. Modigliani on Sereinu (Sipora), Mentawei Islands. Annali del Museo Civico di Storia Naturale di Genova, Series 2, 34, 613–618.
Boulenger, G.A. (1897) A catalogue of the reptiles and batrachians of Celebes, with special reference to the collections made by Drs. P. & F. Sarasin in 1893–1896. Proceedings of the Zoological Society of London, 65, 193–237.
Boulenger, G.A. (1898) Third report on additions to the lizard collection in the Natural History Museum. Proceedings of the Zoological Society of London, 66, 912–923.
http://dx.doi.org/10.1111/j.1096-3642.1898.tb03194.xBoulenger, G.A. (1900) Descriptions of new reptiles and Batrachians from Borneo. Proceedings of the Zoological Society of London, 69, 182–187.
http://dx.doi.org/10.1111/j.1096-3642.1890.tb01716.xBrongersma, L.D. (1942) Notes on scincid lizards. Zoologische Mededelingen, 24, 125–152.
Bryan, J.E., Shearman, P.L., Asner, G.P., Knapp, D.E., Aoro, G. & Lokes, B. (2013) Extreme differences in forest degradation in Borneo: comparing practices in Sarawak, Sabah, and Brunei. PLoS ONE, 8, e69679.
http://dx.doi.org/10.1371/journal.pone.0069679Cannon, C.H., Morley, R.J. & Bush, A.B.G. (2009) The current refugial rainforests of Sundaland are unrepresentative of their biogeographic past and highly vulnerable to disturbance. Proceedings of the National Academy of Sciences of the United States of America, 106, 11188–11193.
http://dx.doi.org/10.1073/pnas.0809865106Cochran, D.M. (1927) New reptiles and batrachians collected by Dr. Hugh M. Smith in Siam. Proceedings of the Biological Society of Washington, 40, 179–192.
Darevsky, I.S. & Orlov, N.L. (2005) New species of limb-reduced lygosomine skink genus Leptoseps Greer, 1997 (Sauria, Scincidae) from Vietnam. Russian Journal of Herpetology, 12, 65–68.
Datta-Roy, A., Das, I., Bauer, A.M., Tron, R.K.L. & Karanth, P. (2013) Lizard wears shades. A spectacled Sphenomorphus (Squamata: Scincidae), from the sacred forests of Mawphlang, Meghalaya, north-east India. Zootaxa, 3701 (2), 257–276.
http://dx.doi.org/10.11646/zootaxa.3701.2.7de Queiroz, K. (1998) The general lineage concept of species. Species criteria, and the process of speciation. In: Howard, D.J. & Berlocher, S.H. (Eds.), Endless forms: Species and Speciation. Oxford University Press, New York, USA, pp. 57–75.
de Queiroz, K. (1999) The general lineage concept of species and the defining properties of the species category. In: Wilson, R.A. (Ed.), Species: New Interdisciplinary Essays. Massachusetts Institute of Technology Press, Cambridge, Massachusetts, pp. 49–89.
Dunn, E.R. (1927) Results of the Douglas Burden Expedition to the island of Komodo. III.—Lizards from the East Indies. American Museum Novitates, 288, 1–13.
Greer, A.E., David, P. & Teynié, A. (2006) The southeast Asian Scincid lizard Siaphos tridigitus Bourret, 1939 (Reptilia, Scincidae): a second specimen. Zoosystema, 28, 785–790.
Grismer, L.L. (2006) Two new species of skinks (genus Sphenomorphus Fitzinger 1843) from the Seribuat Archipelago, West Malaysia. Herpetological Natural History, 9, 151–162.
Grismer, L.L. (2007) A new species of small montane forest floor skink (Genus Sphenomorphus Fitzinger 1843) from southern peninsular Malaysia. Herpetologica, 63, 544–551.
http://dx.doi.org/10.1655/0018-0831(2007)63%5B544:ANSOSM%5D2.0.CO;2Grismer, L.L. (2008) A new species of insular skink (Genus Sphenomorphus Fitzinger 1843) from the Langkawi Archipelago, Kedah, west Malaysia with the first report of the her- petofauna of Pulau Singa Besar and an updated checklist of the herpetofauna of Pulau Langkawi. Zootaxa, 1691, 53–26.
Grismer, L.L. (2015) Paleoendemism and genetic uniformity across 17 different clades in the sky island archipelago of Malaysia. Oral Presentation. Annual Meeting of the Society for the Study of Reptiles and Amphibians, University of Kansas, Lawrence, Kansas.
Grismer, L.L., Ahmad, N. & Onn, C.K. (2009a) A new, diminutive, upland Sphenomorphus Fitzinger 1843 (Squamata: Scincidae) from the Belum-Temengor Forest Complex, Peninsular Malaysia. Zootaxa, 2312, 27–38.
Grismer, L.L., Wood, P.L. & Grismer, J.L. (2009b) A new insular species of skink of the genus Sphenomorphus Strauch 1887 (Squamata: Scincidae) from Pulau Perhentian Besar, Terengganu, Peninsular Malaysia. Tropical Life Sciences Research, 20, 51–69.
Grismer, L.L. & Quah, E.S.H. (2015) The rediscovery of Sphenomorphus malayanus Doria, 1888 (Squamata: Scincidae) from the Titiwangsa Mountain Range of Peninsular Malaysia and its re-description as S. senja sp. nov. Zootaxa, 3931 (1), 63–70.
http://dx.doi.org/10.11646/zootaxa.3931.1.4Inger, R.F., Lian, T.F., Lakim, M. & Yambun, P. (2001) New species of the lizard genus Sphenomorphus, (Lacertilia : Scincidae), with notes on ecological and geographic distribution of species in Sabah, Malaysia. The Raffles Bulletin of Zoology, 49, 181–189.
Kopstein, F. (1926) Reptilien von den Molukken und den benachbarten Inseln. Zoologische Mededelingen, 9, 71–112.
Leaché, A.D. (2009) Species tree discordance traces to phylogeographic clade boundaries in North American fence lizards (Sceloporus). Systematic Biology, 58, 547–559.
http://dx.doi.org/10.1093/sysbio/syp057Linkem, C.W. (2013) Molecular Phylogenetics and Biogeography of Sphenomorphini (Squamata: Scincidae). PhD Thesis, Department of Ecology and Evolutionary Biology, University of Kansas, Lawrence, Kansas. 250 pp.
Linkem, C.W., Diesmos, A.C. & Brown, R.M. (2010) A new species of scincid lizard (genus Sphenomorphus) from Palawan Island, Philippines. Herpetologica, 66, 67–79.
http://dx.doi.org/10.1655/08-074.1Linkem, C.W., Diesmos, A.C. & Brown, R.M. (2011) Molecular systematics of the Philippine forest skinks (Squamata: Scincidae: Sphenomorphus): testing morphological hypotheses of interspecific relationships. Zoological Journal of the Linnean Society, 163, 1217–1243.
http://dx.doi.org/10.1111/j.1096-3642.2011.00747.xMacey, J.R., Larson, A., Ananjeva, N.B., Fang, Z. & Papenfuss, T.J. (1997) Two novel gene orders and the role of light-strand replication in rearrangement of the vertebrate mitochondrial genome. Molecular Biology and Evolution, 14, 91–104.
http://dx.doi.org/10.1093/oxfordjournals.molbev.a025706Müller, F. (1895) Reptilien und amphibien aus Celebes. Verhandlungen der Naturforschenden Gesellschaft in Basel, 10, 825–843.
Nguyen, T.,Q., Schmitz, A., Nguyen, T.T., Orlov, N.L., Böhme, W. & Ziegler, T. (2011) Review of the genus Sphenomorphus Fitzinger, 1843 (Squamata: Sauria: Scincidae) in Vietnam, with description of a new species from northern Vietnam and southern China and the first record of Sphenomorphus mimicus Taylor, 1962 from Vietnam. Journal of Herpetology, 45, 145–154.
http://dx.doi.org/10.1670/09-068.1Rohland, N. & Reich, D. (2012) Cost-effective, high-throughput DNA sequencing libraries for multiplexed target capture. Genome Research, 22, 939–946.
http://dx.doi.org/10.1101/gr.128124.111Stamatakis, A. (2006) RAxML-VI-HPC: maximum likelihood-based phylogenetic analyses with thousands of taxa and mixed models. Bioinformatics, 22, 2688–2690.
http://dx.doi.org/10.1093/bioinformatics/btl446Stejneger, L. (1908) Three new species of lizards from the Philippine Islands. Proceedings of the United States National Museum, 34, 199–204.
http://dx.doi.org/10.5479/si.00963801.34-1606.199Taylor, E.H. (1918) Reptiles of Sulu Archipelago. Philippine Journal of Science, 13, 233–265.
Taylor, E.H. (1962) New Oriental reptiles. University of Kansas Science Bulletin, 43, 209–263.
http://dx.doi.org/10.5962/bhl.part.13346Wiley, E.O. (1978) The evolutionary species concept reconsidered. Systematic Biology, 27, 17–26.
http://dx.doi.org/10.2307/2412809